taxonID	type	description	language	source
03E787F8EB28FFAD5DF0F96BFD003845.taxon	type_taxon	(description, type species: Termitozophilus laetus Silvestri, 1901);	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB28FFAD5DF0F96BFD003845.taxon	discussion	Termitozophilus has been recently redescribed by Zilberman (2019). However, because the four new species revealed in this study more than doubled the number of known species in the genus, a redescription and additional discussion are required.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB28FFAD5DF0F96BFD003845.taxon	description	Head elongate to subquadrate; gula subtriangular, widest at base, fused to mentum; foramen magnum short with vertex bearing four to six bristles and with a strong V-shaped impression in front, between the eye. Antenna with 11 antennomeres, antennomeres shape varying among species, interarticular membrane between antennomeres II and III largely exposed. Labrum more than twice wider than long; five main bristles on each lateral half, distributed in two pairs in diagonal rows and one with variable position. Mandible symmetrical, internal margin with a mesal broad tooth; with pores on surface of mesal region. Maxilla with stipe almost as wide as long, a long bristle at apex, close to first palpomere, and four short bristles close to galea; lacina subquadrate to elongate, covered with bristles of different length at margins; galea narrow, about as long as lacina; surface covered with some moderately long bristles; maxillary palpus densely setose and with four palpomeres with different degree of sclerotization: palpomere I short and subquadrate, palpomeres II – IV elongate; only palpomeres II and III covered with long bristles; palpomeres I – III porose and with a scaled surface, gradually increasing from I to III; palpomere IV not scaled and with sensilla. Labium with epipharynx covered with pores and with short bristles; lateral lobes of epipharynx developed, with a row of bristles on each side; labial palpus setose, 3 - articulated; palpomeres I – II subquadrate; last palpomere elongate and narrow; ligula narrowed at base, with rounded or bifid apex, almost variating in shape among species; mentum and submentum distinct or not; preapical and proximal bristles always present and apical bristle present or not. Abdomen. Inner and outer paratergites fused or not to a single sclerite by secondary sclerotization; paratergites fused or not to tergites or sternites; each inner and outer paratergite with two rows of three to six bristles; tergite VII from subquadrate to about twice as wide as long, with or without bristles; when present, distributed in two rows of long bristles, medial region with two bristles and posterior region with six bristles, short bristles covering entire surface, glandular structure present, continuous along its length (not separated). Aedeagus. Paramere welldeveloped, with two to four bristles on apical lobe and velar sac well developed extending to apical lobe; median lobe with rounded bulbus and with subquadrate tubus apex, with sclerites of the internal sac well developed, variable among species. Spermatheca. Capsule large and sclerotized; chamber and duct fused, curved, with duct placed in posterior region, weakly sclerotized. Oviparity. Knowledge about the reproductive biology of the species in the tribe Corotocini or even in other termitophilous taxa of Staphylinidae is poorly known. Emerson (1935) described a curious case of a Nasutitermes guyanae (Holmgren) nest with four species of Corotocini: Thyreoxenus major Mann, Eburniola leucogaster Mann (both Corotocina), Termitophya amica Mann (Abrotelina) and Xenopelta cornuta Mann (Termitogastrina), where two kinds of larvae were found, as well as three different kinds of eggs, often found among termite eggs. One of the egg kinds was elongate (supposedly belonging to T. major), and the other two were less elongate, with one kind being even smaller than a termite egg, which Emerson (1935) supposedly attributed to E. leucogaster specimens. Eight spherical eggs were found in a dissected female abdomen of Termitozophillus laetus housed at MZSUP (Figs. 2 A – D). Although no Termitozophilus species eggs or larvae have been found in termite nests, we presume this genus is oviparous based on the number and size of eggs reported in T. laetus. The most complete reproductive biology data about a Corotocini species is now available for viviparous species of the genus Corotoca Schiødte, in which females carry three very large eggs in the abdomen with embryos that grow asynchronously (Zilberman et al. 2019). Unlike Termitozophillus, spermatheca of Corotoca is reduced, though the relationship between spermatheca loss or reduction and viviparity, if any, is unclear (Oliveira et al. 2018; Pisno et al. 2019; Zilberman et al. 2019). Although Corotocini biology is little understood, oviparity has been the most documented strategy in genera of this tribe (Emerson 1935, Seevers 1957, Zilberman et al., 2019).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB28FFAD5DF0F96BFD003845.taxon	distribution	Geographic distribution and host relationship. The genus Termitozophilus is found only in the Neotropical Region, from the northernmost areas of South America (Guyana) to Argentina and Paraguay (Zilberman 2019). This distribution is directly related to the distribution of termite hosts (Cornitermes). The genus Cornitermes contains 14 valid species, although only six of them have been shown to harbor Termitozophilus species. As a result, new Termitozophilus species are expected as nests of Cornitermes species that have not been screened for termitophilous beetles are studied. Termitozophilus laetus is the most widely distributed species, appearing in a variety of biomes, including the Amazon Rainforest, where it is associated with C. incisus Emerson, and the Atlantic Forest, Cerrado (savanna-like vegetation), and southern grasslands, where it is associated with C. cumulans (Kollar), C. snyderi Emerson, C. bequaerti Emerson and Cornitermes “ near villosus ”. The polyxenic condition of T. laetus is likely one of the key reasons for its widespread dispersion. Only the type series collected in the Amazon of Guyana, Termitozophilus mirandus which is associated with C. pugnax and a widely distributed species in the Amazon region, is known. In this sense, as it occurs with its termite host, T. mirandus may be widely distributed in the Amazon. Termitozophilus belleae is only known from the type series collected in the Brazilian Cerrado (savanna-like vegetation) in the state of Mato Grosso, in C. silvestri nests. It is, however, a widely distributed host, found across the Cerrado biome and sections of the Brazilian Amazon Rainforest (Constantino, 2002; Zilberman, 2019). Hence, there is a chance that T. belleae distribution will extend, as additional collecting efforts are made in previously unsampled areas. The four species described in this study were found in Cornitermes nests in pastures in the Brazilian Amazon’s northernmost state of Pará.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB28FFAD5DF0F96BFD003845.taxon	discussion	Remarks. The description of four new Termitozophilus species in this study, as well as a review of existing data from descriptions, photographs, and mounted slides of previously reported species, allowed us to expand the generic diagnosis and redescription. Scanning electron microscope images also aided in the advancement of the morphological research by revealing new features. All Termitozophilus species have articular membranes that are largely exposed between antennomeres II-III, as well as pores across the major sclerotized region of sternites. The previously sparse descriptions of the aedeagus and spermatheca are now expanded to include information regarding chaetotaxy of the paramere, velar sac, median lobe, inner sac sclerites (all aspects of male genitalia), and the morphology of the chamber and duct of the spermatheca. In the internal view, the anteromedial border of the elytra is scaled, a condition initially identified in some Termitomorpha Wasmann species (Aleocharinae: Corotocini: Termitogastrina) (Caron et al. 2018). The discovery of the same structure in a separate subtribe emphasizes the need for more inquiry as well as a potential hypothesis of primary homology in a larger sense, maybe extending to other Corotocini tribes. The condition of the paratergites, which can be united to sternites or not, demonstrates significant plasticity in the evolution of abdominal sclerites. The shape and arrangement of abdominal sclerites identified in stenogastric specimens may aid in understanding the evolution of abdominal sclerites. Furthermore, the arrangement of paratergites is also useful for species identification.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	description	(Figs. 3 – 9, 27 B, 28)	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	materials_examined	Type material. HOLOTYPE ♂ (MPEG): Brasil [= Brazil], Pará, Barcarena / Vila de Arapari [= Arapari Village], 05. VIII. 2021 / R. F. Silva [collector], hospedeiro [= host]: Cornitermes sp. PARATYPES. Same data as holotype, 1 ♀ in MPEG (in alcohol), 1 ♀ and 1 ♂ in MZSUP (MZSP 21291) (dissected in slides).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	diagnosis	Diagnosis. Paratergites fused neither to each other nor to tergites and sternites; sclerotized areas of the abdomen light brown, with a large reddish brown membranous area exposed in dorsal view. Head with two pairs of bristles on vertex, one behind eye and one on medial-posterior portion of vertex. Elytra trapezoidal with heavily scaled anteromesial border. Hind wing membrane shed during post-imaginal growth. Sternite VIII with posterior margin bearing two bristles.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	description	Description. Male — length ~ 1.6 mm (Figs. 3 A – B); female — length ~ 1.7 mm (Figs. 4 A – B). Head subquadrate, about as wide as long, widest behind eye, almost narrowing posteriorly; length of eye about 1 / 3 of head length; vertex bearing two pairs of bristles: one bristle behind eye and one bristle on medial-posterior region (Fig. 5 A). Antenna: scape almost longer than remainder antennomeres except antennomere XI; antennomere XI about three times longer than wide and more than twice longer than antennomere X; antennomeres II – VI elongate, gradually decreasing in length, antennomeres VII – X moniliform. Mouthparts. Labrum more than twice wider than long, with slight sinuous anterior margin, five main long bristles on each lateral half, distributed in two diagonal rows each with two bristles and one bristle placed above middle of labrum length (Fig. 5 C); epipharynx covered with sensilla and numerous short bristles, two of which are rather longer. Mandible apex narrow, bearing a tooth on mesal region; with many punctuations on basal two thirds (Figs. 5 B, 7 A). Labium with ligula narrowed at base and with a median-apical emargination, resulting in two lobes; mentum and submentum distinct, both trapezoidal, with three bristles distributed on each lateral margin, as follows: apical, pre-apical, and proximal (Fig. 5 E). Thorax. Elytra almost trapezoidal, wider at posterior region (Fig. 6 C), covering mesonotum and metanotum; anteromesial border densely scaled, in internal view (Figs. 8 A – B); covered with long bristles. Hind wing represented by a small pad (Fig. 6 B). Abdomen with short tergites and with secondary sclerotization, not reaching inner paratergites; outer paratergites close to margin of sternites by the former secondary sclerotization; inner and outer paratergites not fused (Fig. 27 B); first segment represented by a weak sclerotized tergite joined to metanotum (Fig. 6 B); sternites III – V with sparsely distributed pores; tergite VII (Fig. 6 D) almost subquadrate, wider than long, with two rows of bristles: two bristles on medial region and six bristles on posterior region; anterior margin with conspicuous glandular area (Figs. 8 C – D); tergite VIII (Fig. 6 G) subquadrate, with two rows of six long main bristles on medial and apical regions, and four moderately sized bristles on posterior margin; sternite VIII (Fig. 6 H) almost subquadrate, with acute posterior margin, with a row of six medial bristles, a row of eight apical bristles, and two moderately sized bristles on posterior margin; male sternite IX elongate and tongue-shaped, with four bristles on apical region (Fig. 6 F); female sternite IX represented by a pair of hemisternites laterally attached to tergite IX (Fig. 6 I); tergite X not bilobed, with three long bristles on each lateral half, two on medial region and two bristles on posterior region.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	etymology	Etymology. The specific epithet is patronymic in honor of the wildlife photographer, researcher and science communicator César Favacho, for having contributed many photos of termitophilous beetles for this study.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	distribution	Geographic distribution. Brazil (Pará) (Fig. 28).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	biology_ecology	Host termites. This species was found in epigeal (above-ground) nests of an unidentified species of Cornitermes, that possibly belongs to an undescribed species. Behavior. The behavior of Termitozophilus favachoi sp. nov. is similar to that described for T. pilosus sp. nov. (Figs. 9 A – D) (see behaviour under T. pilosus sp. nov.).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB2CFFA05DF0FC92FDAC3A35.taxon	discussion	Discussion. Termitozophilus favachoi sp. nov. shares with T. pilosus sp. nov. and T. parapilosus sp. nov. abdominal sclerites and paratergites fused neither to each other nor to tergites and sternites (Figs. 27 A – E). It differs from T. pilosus sp. nov in having four moderately large bristles on posterior margin of tergite VIII (Fig. 6 G) (six in T. pilosus sp. nov.) and two bristles on sternite VIII (Fig. 6 H) (four in T. pilosus sp. nov.). Termitozophilus favachoi sp. nov. can be distinguished from T. parapilosus sp. nov. in having four bristles on the vertex (Fig. 5 A) (six in T. parapilosus sp. nov.) and two moderately sized bristles on the posterior margin of sternite VIII (Fig. 6 H) (six in T. parapilosus sp. nov.). Additionally, the elytra of T. favachoi sp. nov. has an anteromesial border densely scaled in the internal view (Figs. 8 A – B), a feature shared with T. parapilosus sp. nov. In T. pilosus sp. nov. the anteromesial border of elytra is wrinkled (Figs. 17 A – B).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	description	(Figs. 10 – 13, 27 B, 28)	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	materials_examined	Type material. HOLOTYPE ♂ (MPEG): Brasil [= Brazil], Pará, Peixe-Boi / Fazenda Abalém [= Abalém Farm], 15. VII. 2021 / R. F. Silva [collector], hospedeiro [= host]: Cornitermes pilosus. PARATYPES. Same data as holotype, 1 ♀ in MPEG (in alcohol), 1 ♀ and 1 ♂ in MZSUP (MZSP 21302) (dissected in slides).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	diagnosis	Diagnosis. Paratergites fused neither to each other nor to tergites and sternites; sclerotized parts of the abdomen mostly light brown, with a large white-colored membranous area exposed in dorsal view. Head with three pairs of bristles, one behind the eye, one on medial-posterior region, and one below medial-posterior area. Elytra subquadrate with a heavily scaled anteromesial border. Apex of sternite VIII with six bristles.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	description	Description. Male — length ~ 1.4 mm (Figs. 10 A – B); female — length ~ 1.7 mm (Figs. 10 C – F). Head subquadrate, about as wide as long, widest behind eye, almost narrowing posteriorly; eye about 1 / 3 of head length; three pairs of bristles on vertex, one bristles behind eye, one bristle on medial-posterior region, and one bristle below the medial-posterior region (Fig. 11 A). Antenna: scape almost longer than remainder antennomeres, except antennomere XI; antennomere XI about three times longer than wide and more than twice longer than antennomere X; antennomeres II – VI elongate, gradually decreasing in length, antennomere VII – X moniliform. Mouthparts. Labrum more than twice wider than long, with slight sinuous anterior margin; five main long bristles on each lateral half, arranged in two diagonal rows each with two bristles and one bristle on middle of labrum length (Fig. 11 C); epipharynx covered with sensilla and numerous short bristles, two of which longer. Mandible apex narrow, bearing a tooth on mesal region; with many punctuations on basal 2 / 3 (Fig. 11 B). Labium with ligula narrowed at base and with a median-apical emargination, resulting in two lobes; mentum and submentum distinct, both trapezoidal, with three bristles distributed on each lateral margin, as follows: apical, pre-apical, and proximal (Fig. 11 E). Thorax. Elytra subquadrate, almost longer than wide and almost wider at posterior region (Fig. 12 C), covering mesonotum and metanotum; anteromesial border densely scaled in internal view (Figs. 13 A – B); covered with long bristles. Hind wing membrane shed in post-imaginal stage, present as a stump (Fig. 12 B). Abdomen with short tergites and with secondary sclerotization not reaching inner paratergites; outer paratergites close to margin of sternites by the former secondary sclerotization; inner and outer paratergites not fused (Fig. 27 B); first segment represented by a weak sclerotized tergite joined to metanotum (Fig. 12 B); sternites III – V with pores sparsely distributed; tergite VII (Fig. 12 D) subquadrate, wider than long, with two rows of bristles, two bristles on medial region and six bristles on apical region, anterior margin with conspicuous glandular area; tergite VIII (Fig. 12 G) almost subquadrate, with two rows of bristles, each with six main long bristles on medial and posterior regions, and four moderated sized bristles on posterior margin; sternite VIII (Fig. 12 H) subquadrate, with acute posterior margin; with a row of six medial bristles, a row of eight posterior bristles, and six moderated sized bristles on posterior margin; male sternite IX (Fig. 12 F) elongate and tongue-shaped, with four bristles on posterior half; female sternite IX represented by a pair of hemisternites laterally attached to tergite IX (Fig. 12 I); tergite X (Fig. 12 E) not bilobed, with three long bristles on each side and with two bristles on medial region.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	etymology	Etymology. The specific epithet “ parapilosus ” is a combination of “ para ”, of Greek origin, which means “ close to ” and “ pilosus ”, referring to the species T. pilosus sp. nov., due to the morphological similarity between these species.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	distribution	Geographic distribution. Brazil (Pará).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	biology_ecology	Host termites. This species was found in epigeal (above-ground) nests of Cornitermes pilosus Holmgren. This is the first record of a termitophilous rove beetle in the nest of this termite species.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB22FFA75DF0FB69FAAC38B5.taxon	discussion	Discussion. Termitozophilus parapilosus sp. nov. is similar to T. pilosus sp. nov. and T. favachoi sp. nov. in having paratergites fused neither to each other nor to tergites and sternites (Figs. 27 A – E). Termitozophilus parapilosus sp. nov. can be easily distinguished from T. pilosus sp. nov. in having six bristles on vertex (Fig. 11 A) (T. pilosus sp. nov. has four bristles). In addition, it has four bristles on posterior margin of tergite VIII (Fig. 12 G) (six in T. pilosus sp. nov.) and six bristles on the posterior margin of sternite VIII (Fig. 12 H) (four in T. pilosus sp. nov.). Termitozophilus parapilosus sp. nov. also has anteromesial border of elytra densely scaled in the internal view (Figs. 13 A – B). See the discussion under T. favachoi sp. nov. for differences between these similar new species.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	description	(Figs. 14 – 18, 27 B, 25, 28)	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	materials_examined	Type material. HOLOTYPE ♂ (MPEG): Brasil [= Brazil], Pará, Peixe-Boi / Fazenda Abalém [Abalém Farm], 15. VII. 2021 / R. F. Silva [collector], hospedeiro [= host]: Cornitermes pilosus. PARATYPES. Same data as holotype, 5 ♂♂ and 5 ♀♀ in MZSUP (MZSP 21290, MZSP 21291) (1 ♀ and 1 ♂ dissected in slides, and 8 specimens in alcohol 70 %); 5 ♂♂ and 5 ♀♀ in MPEG (in alcohol 70 %); 1 ♀ and 1 ♂ in the CESP (in alcohol 70 %); and 5 ♂♂ and 5 ♀♀ in the FMNH (in alcohol 70 %).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	diagnosis	Diagnosis. Paratergites fused neither to each other nor to tergites and sternites; sclerotized areas of abdomen mostly light brown, with a large white-colored membranous area exposed in dorsal view. Head with vertex bearing two pairs of bristles, one bristle behind the eye and one on medial-posterior area. Elytra trapezoid with anteromesial border barely scaled. Hind wing membrane shed during post-imaginal growth. Posterior margin of sternite VIII with four bristles.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	description	Description. Male — length ~ 1.3 mm (Figs. 14 C – D); female — length ~ 1.4 mm (Figs. 14 A – B). Head subquadrate, about as wide as long, widest behind eye, almost narrowing posteriorly; eye about 1 / 3 of head length; vertex bearing two pairs of bristles: one bristle behind eye and one bristle on medial-posterior region (Fig. 15 A). Antenna: scape longer than remainder antennomeres, except antennomere XI; antennomere XI about three times longer than wide and more than twice longer than antennomere X; antennomeres II – VI elongate, gradually decreasing in length, antennomeres VII – X moniliform. Mouthparts. Labrum more than twice wider than long, with slight sinuous anterior margin; five main long bristles on each lateral half, arranged in two diagonal rows each with two bristles and one bristle placed above middle of labrum length (Fig. 15 C); epipharynx covered with sensilla and numerous short bristles, two of which longer. Mandible apex narrow, bearing a tooth on mesal region; with many punctuations on basal two thirds (Figs. 15 B, 18 A – C). Labium with ligula narrowed at base and with a median-apical emargination, resulting in two lobes; mentum and submentum distinct, both trapezoidal, with three bristles distributed on each lateral margin, as follows: apical, pre-apical, and proximal (Fig. 17 D). Thorax. Elytra trapezoidal, longer than wide and wider at posterior region (Fig. 16 C), covering mesonotum and metanotum; anteromesial border barely scaled in internal view (Figs. 17 A – B); covered with long bristles. Hind wing membrane shed during post-imaginal growth, present as a stump (Fig. 16 B). Abdomen with short tergites and with secondary sclerotization, not reaching inner paratergites; outer paratergites close to sternites margin by the former secondary sclerotization; inner and outer paratergites not fused (Fig. 27 B); first segment represented by a weak sclerotized tergite joined to metanotum (Fig. 16 B); sternites III – V with sparsely distributed pores; tergite VII (Figs. 16 D, 18 D – E) subquadrate, wider than long, with two rows of bristles, two bristle on medial region and six bristles on posterior half, anterior margin with conspicuous glandular area; tergite VIII (Fig. 16 G) almost subquadrate, with two rows of bristles, each with six main long bristles on medial and posterior regions, and with six moderated sized bristles on posterior margin; sternite VIII (Fig. 16 H) subquadrate, with acute posterior margin, with a row of six medial bristles, a row of eight posterior bristles, and four moderated sized bristles on posterior margin; male sternite IX (Fig. 16 F) elongate and tongue-shaped, with four bristles on posterior half; female sternite IX represented by a pair of hemisternites laterally attached to tergite IX (Fig. 16 I); tergite X (Fig. 16 E) not bilobed, with three long bristles on each lateral half and two bristles on medial region.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	etymology	Etymology. The specific epithet “ pilosus ” referring to the host species “ C. pilosus ”.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	distribution	Geographic distribution. Brazil (Pará).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	biology_ecology	Host termites. This species was found in epigeal (above-ground) nests of Cornitermes pilosus. This is the first record of a termitophilous rove beetle in the nest of this termite species. Notes on behavior. During laboratory observations (Fig. 1 F), T. pilosus sp. nov. appears to be completely integrated into the colony of its host termite, since no agonistic interactions were observed, and the termites were seen palpating the beetles’ antennae, mouthparts, and abdomens. This species spent much of its time with the distal region of abdomen curled up over its body, not leaving the dorsal portion of the abdomen exposed. However, on other occasions, the abdomen was extended, exposing the tergites, which attracted termites that licked the gland present on tergite VII.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB39FFBD5DF0FF5EFEE63D7D.taxon	discussion	Discussion. Termitozophilus pilosus sp. nov. is similar to T. parapilosus sp. nov. in having abdominal sclerites mostly light brown and paratergites fused neither to each other nor to tergites and sternites (Figs. 27 A – E). See discussion under T. favachoi sp. nov. and T. parapilosus sp. nov. for the resemblances and differences between these species.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	description	(Figs. 19 – 24, 27 E, 28)	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	materials_examined	Type material. HOLOTYPE ♂ (MPEG): Brasil [= Brazil], Pará, Magalhães Barata / Vila de Calafate [= Calafate Village], 11. III. 2021 / R. F. Silva [collector] / hospedeiro [= host]: Cornitermes pilosus. PARATYPES. Same data as holotype, 5 ♀♀ and 5 ♂♂ in MPEG (in alcohol 70 %), 1 ♀ and 1 ♂ in MZSUP (MZSP 21288) (dissected on slides).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	diagnosis	Diagnosis. Paratergites fused neither to each other nor to tergites and sternites; sclerotized areas mostly dark brown, with a large white membranous area exposed in dorsal view. Head with vertex bearing two pairs of bristles, one behind the eye and a pair on medial-posterior region. Hind wings present in post-imaginal growth. Elytra trapezoidal, with anteromesial border densely scaled. Posterior margin of sternite VIII with two bristles.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	description	Description. Male — length ~ 1.6 mm (Figs. 19 A – B); female — length ~ 1.8 mm (Figs. 20 A – C). Head subquadrate, about as wide as long, widest behind eye, narrowing conspicuously through posterior region; eye about 1 / 3 of head length; vertex bearing two pairs of bristles: one bristle behind eye and one on medialposterior region (Fig. 21 A). Antenna: scape almost longer than remainder antennomeres, except antennomere XI; antennomere XI about three times longer than wide and more than twice longer than X; antennomeres II – VI elongate, gradually decreasing in length, VII – X moniliform. Mouthparts. Labrum more than twice wider than long, with almost sinuous anterior region; five main long bristles on each lateral half, distributed in two diagonal rows, each with two bristles and one bristle placed on middle of labrum length (Fig. 21 C); epipharynx covered with sensilla and numerous short bristles, two of which longer. Mandible apex narrow, bearing a tooth on mesal region; with many punctuations on basal two thirds, and with some pores on outer margin (Figs. 21 B, 23 C – D). Labium with ligula narrowed at base and with a median-apical emargination, resulting in two lobes; mentum and submentum distinct, both trapezoidal, with three bristles distributed on each lateral margin, as follows: apical, pre-apical, and proximal (Figs. 21 E, 24 C). Thorax. Elytra trapezoidal, almost longer than wide and wider at posterior region, covering mesonotum and metanotum (Fig. 22 B); anteromesial border densely scaled in internal view (Figs. 23 A – B); covered with long bristles. Hind wing present, membrane not shed during post-imaginal growth (Fig. 22 A). Abdomen with short tergites and with secondary sclerotization not reaching inner paratergites; outer paratergites fused to sternites by former secondary sclerotization; inner and outer paratergites fused (Fig. 27 E); first segment represented by a strong sclerotized tergite attached to metanotum, with medial region strongly projected backwards (Fig. 22 A); sternites III – V with pores clustered; tergite VII (Fig. 22 C) almost subquadrate, wider than long, with two rows of bristles, two bristles on medial region and six bristles on posterior half, anterior margin of tergite VII with conspicuous glandular areas (Figs. 24 E – F); tergite VIII (Fig. 22 F) almost subquadrate, with two rows of bristles, six main long bristles on medial and posterior regions; sternite VIII (Fig. 22 G) subquadrate, with one pair of median bristle, one row of six bristle close to posterior margin and eight bristles along posterior margin; male sternite IX (Fig. 22 E) elongate and tongue-shaped, with two bristles on posterior half; female sternite IX (Fig. 22 H) represented by a pair of hemisternites laterally attached to tergite IX; tergite X (Fig. 22 D) not bilobed, with three long bristles on each lateral half, two on medial region and two on posterior region.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	etymology	Etymology. The specific epithet is a Latin adjective that means “ dark ”, alluding to dull coloration of the examined specimens.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	distribution	Geographic distribution. Brazil (Pará).	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
03E787F8EB3CFFB45DF0F9BAFB843931.taxon	discussion	Discussion. Termitozophilus tenebrus sp. nov. is similar to T. laetus in having inner and outer paratergites fused, and paratergites fused to sternites (Figs. 27 A – E). It differs from T. laetus in having head, thorax and abdomen with dark brown sclerites (light brown in T. laetus) and pores on sternites distributed in circular clusters (Figs. 19 A – B, 20 A – C) (scattered in T. laetus). In addition, T. tenebrus sp. nov. has a trapezoidal elytron (Fig. 22 B), tergite VII with bristles (Fig. 22 C) and posterior margin of tergite VIII without bristles (Fig. 22 F). In T. laetus, the elytron is subquadrate, the bristles on tergite VII are absent and the posterior margin of tergite VIII has four bristles (see figs. 16 – 18 in Zilberman (2019 )). Termitozophilus tenebrus sp. nov. is associated with nests of C. pilosus in pastures, while T. laetus is associated with nests of C. cumulans in savanna-like environments.	en	Silva, Ruan Felipe Da, Zilberman, Bruno, Carvalho-Filho, Fernando Da Silva (2022): Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 5209 (5): 501-534, DOI: 10.11646/zootaxa.5209.5.1
